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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
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<publisher-name>Frontiers Media S.A.</publisher-name>
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<article-id pub-id-type="doi">10.3389/fpls.2024.1354672</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>A decade of advances in the study of buckwheat for organic farming and agroecology (2013-2023)</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Vieites-&#xc1;lvarez</surname>
<given-names>Yedra</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
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<xref ref-type="aff" rid="aff2">
<sup>2</sup>
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<contrib contrib-type="author">
<name>
<surname>Reigosa</surname>
<given-names>Manuel J.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
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<contrib contrib-type="author" corresp="yes">
<name>
<surname>S&#xe1;nchez-Moreiras</surname>
<given-names>Adela M.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
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<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
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<aff id="aff1">
<sup>1</sup>
<institution>Universidade de Vigo, Departamento de Biolox&#xed;a Vexetal e Ciencias do Solo, Facultade de Biolox&#xed;a</institution>, <addr-line>Vigo</addr-line>, <country>Spain</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Instituto de Agroecolox&#xed;a e Alimentaci&#xf3;n (IAA), Universidade de Vigo</institution>, <addr-line>Ourense</addr-line>, <country>Spain</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Daniela Grulova, University of Pre&#x161;ov, Slovakia</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Vera Raji&#x10d;i&#x107;, University of Ni&#x161;, Serbia</p>
<p>Jozef Fej&#xe9;r, University of Pre&#x161;ov, Slovakia</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Adela M. S&#xe1;nchez-Moreiras, <email xlink:href="mailto:adela@uvigo.gal">adela@uvigo.gal</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>06</day>
<month>03</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>15</volume>
<elocation-id>1354672</elocation-id>
<history>
<date date-type="received">
<day>12</day>
<month>12</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>12</day>
<month>02</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Vieites-&#xc1;lvarez, Reigosa and S&#xe1;nchez-Moreiras</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Vieites-&#xc1;lvarez, Reigosa and S&#xe1;nchez-Moreiras</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>During the last decade, research has shown the environment and human health benefits of growing buckwheat (<italic>Fagopyrum</italic> spp.). This comprehensive review aims to summarize the major advancements made in the study of buckwheat from 2013 to 2023, focusing on its agronomic characteristics, nutritional value, and potential applications in sustainable agriculture. The review examines the diverse applications of buckwheat in organic and agroecological farming systems, and discusses the ability of buckwheat to control weeds through allelopathy, competition, and other sustainable farming methods, such as crop rotation, intercropping and green manure, while improving soil health and biodiversity. The review also explores the nutritional value of buckwheat. It delves into the composition of buckwheat grains, emphasizing their high protein content, and the presence of essential amino acids and valuable micronutrients, which is linked to health benefits such as lowering cholesterol levels, controlling diabetes and acting against different types of cancer, among others. Finally, the review concludes by highlighting the gaps in current knowledge, and proposing future research directions to further optimize buckwheat production in organic or agroecological farming systems. It emphasizes the need for interdisciplinary collaboration, and the integration of traditional knowledge with modern scientific approaches to unlock the full potential of buckwheat as a sustainable crop.</p>
</abstract>
<kwd-group>
<kwd>buckwheat</kwd>
<kwd>organic farming</kwd>
<kwd>agroecology</kwd>
<kwd>polyphenols</kwd>
<kwd>sustainable weed management</kwd>
</kwd-group>
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<ref-count count="249"/>
<page-count count="20"/>
<word-count count="11181"/>
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<meta-value>Crop and Product Physiology</meta-value>
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</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>The existence of a crop with allelopathic activity, strong competitive traits against weeds, high protein, vitamin and fiber contents, and antioxidant, anticancer, antihypertension, antibacterial and anti-inflammatory potential sounds unreal, but it is not, is just buckwheat. An emergent functional crop.</p>
<sec id="s1_1">
<label>1.1</label>
<title>Buckwheat</title>
<p>Buckwheat (<italic>Fagopyrum</italic> sp.) is an herbaceous crop belonging to the family Polygonaceaea, whose name comes from the triangular shape of buckwheat seeds. Buckwheat is one of the most unknown alternative gluten-free pseudo-cereal crops originated in the southwest of China, around the mid-6<sup>th</sup> millennium BC, and spread to Europe from around the 3<sup>rd</sup> millennium (<xref ref-type="bibr" rid="B97">Hunt et&#xa0;al., 2018</xref>). Buckwheat seeds look very similar to a grain, but they are not grains, that&#x2019;s why buckwheat does not belong to the family Gramineae or Poaceae (<xref ref-type="bibr" rid="B70">Fletcher, 2016</xref>). There are 15 species described belonging to the <italic>Fagopyrum</italic> genus, but the two most worldwide cultivated species are the common buckwheat (<italic>Fagopyrum esculentum</italic> Moench.) and the Tartary buckwheat (<italic>Fagopyrum tataricum</italic> Gaertn.) (<xref ref-type="bibr" rid="B59">D&#x17e;afi&#x107; and &#x17d;uljevi&#x107;, 2022</xref>). The morphology of the achene is one of the most important differences between <italic>F. tataricum</italic> and <italic>F. esculentum</italic>. <italic>F. tataricum</italic> achenes are grooved with angles that are rounded in the bottom and sharply acute in the top, whereas <italic>F. esculentum</italic> achenes are not grooved with sharply acute angles (<xref ref-type="bibr" rid="B176">Rangappa et&#xa0;al., 2023</xref>). These differences also influence their ability to acclimatize to cold and drought conditions. Morphologically, the stem is usually round and hollow, and frequently changes from green to red. The flowers are usually a cymose cluster that needs to be pollinated to produce seeds. Buckwheat has a long flowering period and serves as a source of nectar for honey. The root system is dense and fibrous, and has a deep main root. Most of the root system is concentrated in the first 25&#xa0;cm of the soil (<xref ref-type="bibr" rid="B176">Rangappa et&#xa0;al., 2023</xref>). Buckwheat has a short life cycle, low nutritional requirements, and abundant biomass production (<xref ref-type="bibr" rid="B167">Pinski et&#xa0;al., 2023</xref>), which make this plant an ideal cover crop in organic or agroecological farming systems.</p>
<p>Regarding the nutritional profile of buckwheat, the seeds are rich in protein (much more than cereals), amino acids, and minerals, showing high contents of lysine, tryptophan, arginine, sterols, vitamins, and phenolic compounds (PC). These compounds (PC) are the most important molecules in buckwheat, and are responsible for the strong value of this crop in organic farming and pharmaceutical studies. Among these specialized metabolites is remarkable the flavonoid rutin, which has been reported to show cardioprotective, vasoprotective, antihypertensive, anti-inflammatory, cytoprotective, and anti-diabetic properties (<xref ref-type="bibr" rid="B39">Chrungoo and Chettry, 2021</xref>).</p>
<p>Surprisingly, although considered an emergent crop, buckwheat production was about 2,263,764.35 tons in 2013, and the cultivation covered 2,263,608 ha worldwide, while just 7 years later buckwheat production decreased to 187,5067.97 tons and the cultivation covered 1,988,534 ha worldwide in 2021 (<ext-link ext-link-type="uri" xlink:href="https://www.fao.org/faostat/en/#data/QCL">https://www.fao.org/faostat/en/#data/QCL</ext-link>), probably related to more than one reason, as discussed by <xref ref-type="bibr" rid="B168">Pirzadah and Rehman (2021)</xref>, who listed a list of potential reasons including, among others, erratic yield, indefinite growing cycle, tendency for abortion of flowers, sensitivity to freezing conditions, or the presence of allergy-inducing compounds, as well as consumers&#x2019; habits and economic limitations. Nonetheless, buckwheat must be still seen as a pseudo-cereal crop with strong potential for functional food sector in the near future.</p>
</sec>
<sec id="s1_2">
<label>1.2</label>
<title>Weeds in buckwheat agroecosystems</title>
<p>The definition of weed is linked to land uses and human interest. Actually, many definitions of weed have been proposed by the scientific community from various perspectives: agronomic, biological, ecological, etc. The most commonly used definitions have been those provided by the European Weed Research Society in 1986 (&#x201c;any plant or vegetation, excluding fungi, interfering with the objectives or requirements of people&#x201d;) and the Weed Science Society of America in 1989 (&#x201c;a plant growing where it is not desired&#x201d;) (<xref ref-type="bibr" rid="B188">Scavo and Mauromicale, 2020</xref>). Weeds are generally considered annoying, aggressive and competitive. Not only due to the weed-related high yield loses but also by their direct impact on the dietary quality of food, hosting insects, pests and other crop pathogens, decreasing land value and interference with water management (<xref ref-type="bibr" rid="B248">Zimdahl, 2018</xref>; <xref ref-type="bibr" rid="B188">Scavo and Mauromicale, 2020</xref>).</p>
<p>At the time when these definitions were established, synthetic herbicides began to be used more assiduously with fast and high effectiveness, eliminating weeds, and preventing the loss of crop productivity (<xref ref-type="bibr" rid="B101">Jabran et&#xa0;al., 2015</xref>). Unfortunately, due to the massive and indiscriminate use of synthetic herbicides over the last years, many weed species became resistant, avoiding the mode of action of these synthetic chemical compounds and resulting in an important problem, which needs to be urgently resolved. During the last decade, the study of weed resistance has been the focus of several agronomic research studies. The increase of this resistance development has been so fast that while (<xref ref-type="bibr" rid="B91">Heap, 2014</xref>) reported, in the international survey of herbicide resistant weeds, 220 herbicide-resistant weeds in 2013, with 404 unique cases (species x site of action) globally, only ten years later, in (<xref ref-type="bibr" rid="B90">Heap, 2023</xref>), the number of herbicide-resistant species increased up to 267 with 513 unique cases. Moreover, more than 20% of weeds were found to show multiple resistance, e.g., resistance to different herbicide groups in the same biotype, which complicates even more the control of these species in the agroecosystems.</p>
<p>Regarding buckwheat, the most problematic weed species can vary depending on the region and the specific growing conditions, but there are some of them that are very extended all over Europe, and most of them have showed resistance to some type of synthetic herbicide group. This is the case of <italic>Amaranthus retroflexus</italic> L., which is one of the most harmful weed species to buckwheat and has been reported to show resistance to the synthetic herbicides metamitron (electron transport inhibitor) and fomesafen (protoporphyrinogen oxidase (PPO) enzyme inhibitor) (<xref ref-type="bibr" rid="B2">Adamczewski et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B94">Huang et&#xa0;al., 2020</xref>); <italic>Chenopodium album</italic> L., which has presented also resistance to metamitron; <italic>Echinochloa crus-galli</italic> (L.) Beauv. that is resistant to acetolactate synthase (ALS) inhibitors (34 cases), acetyl-CoA carboxylase (ACCase) inhibitors (23 cases), photosystem-II inhibitors (11 cases), auxin mimics/cellulose biosynthesis inhibitors (9 cases), very long chain fatty acid inhibitors (6 cases), and microtubule assembly inhibitors (1 case) (<xref ref-type="bibr" rid="B45">Damalas and Koutroubas, 2023</xref>); <italic>Bidens pilosa</italic> L., which is resistant to photosystem II (PSII) and ALS inhibitor herbicides such us imazethapyr and atrazine (<xref ref-type="bibr" rid="B216">Takano et&#xa0;al., 2016</xref>); <italic>Lolium rigidum</italic> Gaud., which showed resistance to acetyl-CoA carboxylase (ACCase), acetolactate synthase (ASL), photosystem II (PSII), 5-enolpyruvylshikimate-3-phosphate synthase (EPSPS), glutamine synthase, very long-chain fatty acid (VLCFA) synthesis, and protoporphyrinogen oxidase (PPO) inhibiting herbicides (<xref ref-type="bibr" rid="B68">Fern&#xe1;ndez-Moreno et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B138">Loureiro et&#xa0;al., 2017</xref>); and <italic>Portulaca oleracea</italic> L., which developed resistance to PSII inhibitors (<xref ref-type="bibr" rid="B163">Okafor et&#xa0;al., 2014</xref>).</p>
<p>However, a plant species only becomes a weed if the cultivation conditions and the management of the agroecosystem favor the spread of this species. In fact, in a multi-diverse and balanced agroecosystem, plant species would behave as spontaneous flora, promoting biodiversity and crop development, not behaving as weeds (<xref ref-type="bibr" rid="B248">Zimdahl, 2018</xref>). <xref ref-type="bibr" rid="B93">Hu et&#xa0;al. (2023)</xref>, analyzed in a review the 100 essential questions about the future of agriculture, and identified four Key points to understand the agricultural systems in a holistic way: (i) Resource and Environment, (ii) Agricultural Production, (iii) Nutrition and Health, and (iv) Social and Economic Impacts. These perspectives support that agriculture has a substantial impact on environment, human health, and society, while producing food and other essential goods (such as fiber, biofuels, medicine, etc.). Agriculture relies on the natural resources, which must be sustainably managed to ensure productive and healthy farming systems for long-term viability. Moreover, agricultural production should be effectively and sustainably conducted to maximize yield, while minimizing adverse environmental effects. Agriculture&#x2019;s main goals are nutrition and health, which include social and economic aspects, as well as the health of humans and the planet (<xref ref-type="bibr" rid="B93">Hu et&#xa0;al., 2023</xref>). Therefore, the tomorrow&#x2019;s agriculture should focus on increasing biodiversity in the agricultural soils (also plant biodiversity), and favoring an autonomous healthy system that can manage the occurrence of a diversity of plants species in the crop fields without damaging crop plants or reducing their yield. Hence, this review proposes some tested and effective agroecological/organic methods to manage spontaneous plant species without the use of herbicides. Alternative weed management strategies will imply a change of paradigm in the current agriculture, implementing a holistic approach that keeps an eye on the whole ecosystem instead of on the plant alone (<xref ref-type="bibr" rid="B86">Gonz&#xe1;lez-Andujar, 2023</xref>).</p>
</sec>
<sec id="s1_3">
<label>1.3</label>
<title>Agroecological strategies</title>
<p>Agroecology (AE), which emerged almost a century ago, is the application of ecological science to the study and design of sustainable agriculture (<xref ref-type="bibr" rid="B13">Altieri, 1995</xref>). AE will allow farmers to increase the power and control of their production while minimizing social and ecological costs from agriculture, such as soil degradation, water contamination, exhaustion of non-renewable resources, susceptibility to climate change, and inequitable social structures (<xref ref-type="bibr" rid="B149">Mier y Ter&#xe1;n Gim&#xe9;nez Cacho et&#xa0;al., 2018</xref>). AE has gained popularity in recent years by appearing as the most comprehensive solution to the many challenges confronting the agri-food systems (<xref ref-type="bibr" rid="B84">Gliessman and Montenegro, 2021</xref>). It is regarded as the bedrock for transforming the conventional agri-food systems to more sustainable and resilient agroecosystems, mitigating the environmental impacts on agriculture while ensuring efficiency in the use of natural resources (<xref ref-type="bibr" rid="B83">Gliessman, 2021</xref>). AE seeks to integrate (agro) ecology science with agricultural practice and social concerns (<xref ref-type="bibr" rid="B238">Wezel et&#xa0;al., 2020</xref>) for the development of sustainable, productive, and resilient agroecosystems.</p>
<p>Different solutions based on agroecological concepts, principles and practices have been developed to preserve productivity and food security in the long-term, while providing ecological benefits and reducing negative external impacts (<xref ref-type="bibr" rid="B66">FAO, 2019</xref>; <xref ref-type="bibr" rid="B18">Atta-Krah et&#xa0;al., 2021</xref>). Practices carried out in agroecological systems include the selection of crops with allelopathic potential, crops with strong competitive traits against weeds, cover crops, mulching, crop rotation, intercropping, etc.</p>
<p>On the other hand, according to the <xref ref-type="bibr" rid="B4">Agricultural and Rural Development (2023)</xref>, organic farming aims to produce food using natural preparations and processes. Organic farming has a limited environmental impact by promoting responsible use of energy and natural resources, maintenance of biodiversity, preservation of regional ecological balance, enhancement of soil fertility and maintenance of water quality. Organic farming regulations in the European Union are intended to provide a clear structure for the production of organic goods throughout the EU, and are designed to satisfy consumer demands for reliable organic products, while providing equal opportunities for producers, distributors, and marketers. According to European Commission&#x2019;s Agriculture and Rural Development, and US-EPA (<xref ref-type="bibr" rid="B225">United States Environmental Protection Agency Integrated pest management (IPM) principles</xref>), some of the practices carried out in an organic farming system would include, similarly to agroecological systems:</p>
<list list-type="simple">
<list-item>
<p>(I) The screening and selection of naturally resistant varieties to pests and diseases (i.e., allelopathic and/or competitive crops against weeds) would reduce the reliance on synthetic pesticides and promote plant health.</p>
</list-item>
<list-item>
<p>(II) Crop rotation. This sustainable farming method is based on growing different crops on the same land in a planned sequence, which will break pest and disease cycles and will take advantage of allelopathic and competitive traits of crops.</p>
</list-item>
<list-item>
<p>(III) Cover crops and green manures: Cover crops are grown to protect and improve the soil, preventing erosion, suppressing weeds, enhancing soil fertility, and providing habitat for beneficial insects and microorganisms. Green manures, which are cover crops specifically grown to be incorporated into the soil, add organic matter and nutrients.</p>
</list-item>
<list-item>
<p>(IV) Composting: is the process of decomposing organic materials, such as plant residues and animal manure, to create nutrient-rich compost. Organic farmers can use compost to enrich the soil, improve its structure, and enhance nutrient availability for plants.</p>
</list-item>
<list-item>
<p>(V) Soil management: the above-mentioned practices are used to enhance soil health, promoting healthy crop growth and enhancing soil fertility and structure.</p>
</list-item>
<list-item>
<p>(VI) Integrated Pest Management (IPM): organic farmers employ IPM strategies to manage pests and diseases. This approach involves a combination of cultural, physical and mechanical practices, biological control methods, and the use of approved organic pesticides when necessary.</p>
</list-item>
</list>
<p>In this context, the goal of this review is synthesizing the advances made in the use of buckwheat for agroecology or organic agriculture systems over the last decade (2013-2023), and providing updated information on the importance and versatility of buckwheat as crop, with the aim of increasing buckwheat production.</p>
</sec>
</sec>
<sec id="s2">
<label>2</label>
<title>Phenolic compounds: occurrence and function</title>
<p>Phenolic compounds, the most ubiquitous compounds in buckwheat, are present in most of the plant organs of this crop, including fruits, seeds, leaves, stems, roots, etc (<xref ref-type="bibr" rid="B50">De la Rosa et&#xa0;al., 2019</xref>). Phenolic compounds are specialized metabolites produced by the phenylpropanoid and shikimic acid pathways, including at least one aromatic ring with one hydroxyl group (<xref ref-type="bibr" rid="B24">Bhatla and Lal, 2018</xref>). Currently, more than 8000 phenolic compounds have been identified so far (<xref ref-type="bibr" rid="B233">Vuolo et&#xa0;al., 2019</xref>). These specialized metabolites can be divided into different groups according to their skeletal structure (i.e., monomeric, dimeric or polymeric phenolics). For example, flavonoids are polymeric phenolics with a C6-C3-C6 skeleton, while phenolic acids are dimeric phenolics with a C6-C1 skeleton. Other phenolic compounds may present other type of basic skeleton as C6-C3 (hydroxycinnamic acid), C6-C2-C6 (stilbenes), (C6&#x2014;C3)n (lignins), etc (<xref ref-type="bibr" rid="B131">Lattanzio, 2013</xref>).</p>
<sec id="s2_1">
<label>2.1</label>
<title>Flavonoids</title>
<p>Among the phenolic compounds, flavonoids are the most abundant in food and are considered to be the most bioactive phenolics in terms of antioxidant, antihypertensive and anti-inflammatory properties (<xref ref-type="bibr" rid="B50">De la Rosa et&#xa0;al., 2019</xref>). The presence of these compounds in buckwheat has been recently reviewed by <xref ref-type="bibr" rid="B146">Matsui and Walker (2020)</xref>, focusing on their synthesis and their presence in various plant organs of buckwheat. Moreover, a recent study has shown the variation of flavonoids composition on <italic>F. esculentum</italic> (common buckwheat, CB) and <italic>F. tataricum</italic> (Tartary buckwheat, TB), two different species of buckwheat, finding that after sowing both species, rutin tended to decrease in stems, leaves and flowers of CB, while tended to increase in leaves and flowers of TB, although decreased amount of rutin could be found in the stems of this buckwheat species. In fact, the maximum peak of rutin was found in TB flowers 50 days after sowing. Quercetin tended to decrease in the stem of CB and all organs of TB, while increasing in leaves and flowers of CB, with the highest peak of quercetin found in TC flowers 40 days after sowing, similarly to rutin. Finally, quercitrin was only found in CB flowers (<xref ref-type="bibr" rid="B116">Kim et&#xa0;al., 2023</xref>).</p>
<p>The main structure of flavonoids includes two phenyl rings (A and B) joined through a heterocyclic pyran ring (C) (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). Depending on the B-ring position and the degree of unsaturation and oxidation of the C-ring, there will be different types of flavonoids; i.e., isoflavones are flavonoids with the B-ring attached to position 3 of the C-ring, while neoflavonoids have the B-ring attached at position 4 or position 2. This last group can be further subdivided into several subgroups based on the structural features of the C-ring: chalcones, anthocyanins, flavones, flavonols, flavanones, and flavanols (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>; <xref ref-type="bibr" rid="B165">Panche et&#xa0;al., 2016</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Basic skeleton of flavonoids formed by two phenyl rings <bold>(A, B)</bold> joined through a heterocyclic pyran ring <bold>(C)</bold>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1354672-g001.tif"/>
</fig>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Flavonoids and other phenolic compounds divided in subclasses. Flavonoids include anthocyanins, chalcone, flavanones, flavones, flavanols and isoflavonoids. Other phenolic compounds are phenolic acids, stilbene, lignans and curcuminoids.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1354672-g002.tif"/>
</fig>
<sec id="s2_1_1">
<label>2.1.1</label>
<title>Isoflavonoids</title>
<p>Isoflavonoids, also known as phytoestrogens, are naturally occurring non-steroidal phenolic plant compounds (<xref ref-type="bibr" rid="B122">K&#x159;&#xed;&#x17e;ov&#xe1; et&#xa0;al., 2019</xref>) with the B-ring attached to the third position of C-ring and mostly found in soybean and legumes (<xref ref-type="bibr" rid="B165">Panche et&#xa0;al., 2016</xref>). Genistein, daidzein, glycitein, biochanin A, and formononetin are the main types of isoflavonoids (<xref ref-type="bibr" rid="B46">Danciu et&#xa0;al., 2018</xref>). These compounds, which have fungistatic, antibacterial, antiviral, and antioxidant properties (<xref ref-type="bibr" rid="B122">K&#x159;&#xed;&#x17e;ov&#xe1; et&#xa0;al., 2019</xref>), act as phytoalexins, antimicrobial compounds found to be accumulated in plants after infections, and help to limit pathogen spread (<xref ref-type="bibr" rid="B5">Ahmed and Kovinich, 2021</xref>). They also have functions in plant-microbe interactions, including defense mechanisms and rhizobia-legume symbiosis (<xref ref-type="bibr" rid="B181">R&#xed;podas et&#xa0;al., 2013</xref>). Ten different isoflavones have been characterized in buckwheat (<xref ref-type="bibr" rid="B137">Li et&#xa0;al., 2019</xref>). However, their weed suppression potential has not been well studied (<xref ref-type="bibr" rid="B202">Sohn et&#xa0;al., 2021</xref>).</p>
</sec>
<sec id="s2_1_2">
<label>2.1.2</label>
<title>Chalcones</title>
<p>The name chalcones derives from the Greek word &#x201c;chalcos&#x201d; and means &#x201c;bronze&#x201d;, which is the color of most of the natural chalcones (<xref ref-type="bibr" rid="B247">Zhuang et&#xa0;al., 2017</xref>). Chalcones are open chain flavonoids with a 15-carbon structure characterized by the absence of a &#x2018;C-ring&#x2019; in the basic flavonoid skeleton structure (<xref ref-type="bibr" rid="B52">D&#xed;az-Tielas et&#xa0;al., 2016</xref>). In terms of human health, chalcone has therapeutic activities including anticancer (<xref ref-type="bibr" rid="B139">Mahapatra et&#xa0;al., 2015</xref>), antioxidant, anti-inflammatory (<xref ref-type="bibr" rid="B88">Guazelli et&#xa0;al., 2021</xref>), antimicrobial (<xref ref-type="bibr" rid="B226">U&#x161;jak et&#xa0;al., 2019</xref>), antiulcer (<xref ref-type="bibr" rid="B187">Sashidhara et&#xa0;al., 2015</xref>), anthelmintic, antiviral and antiprotozoal activities (<xref ref-type="bibr" rid="B87">Goyal et&#xa0;al., 2021</xref>). In plants, chalcones are the precursors of many other phenolic compounds (<xref ref-type="bibr" rid="B182">Rudrapal et&#xa0;al., 2021</xref>), and not only play an important role in antioxidant activity, but also the analogues, <italic>cis</italic>-chalcone and <italic>trans</italic>-chalcone have shown strong phytotoxic potential against the development of adventitious species such as <italic>Plantago lanceolata</italic> L., against the early root growth of <italic>A. retroflexus</italic>, and <italic>E. crus-galli</italic>, and against the development and growth of adult <italic>Arabidopsis thaliana</italic> L. Heynh. plants (<xref ref-type="bibr" rid="B53">D&#xed;az-Tielas et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B38">Chotsaeng et&#xa0;al., 2019</xref>).</p>
<p>In buckwheat, chalcone related enzymes [i.e., chalcone synthase (CHS) and chalcone isomerase (CHI)] are biogenetic precursors of flavonoids and isoflavonoids (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>) (<xref ref-type="bibr" rid="B175">Rammohan et&#xa0;al., 2020</xref>), and some of them, as naringenin chalcone, have been related with important antioxidant roles into the buckwheat plant metabolism (<xref ref-type="bibr" rid="B57">Dumitru et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B95">Huda et&#xa0;al., 2021</xref>). Moreover, CHS gene expression is also increased in buckwheat under stressors like UV light, bacterial or fungal infection, suggesting its antioxidant function (<xref ref-type="bibr" rid="B192">Schenke et&#xa0;al., 2019</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Chalcone as precursor of many flavonoids such as flavone, flavonol, anthocyanidin and isoflavone in plants.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1354672-g003.tif"/>
</fig>
</sec>
<sec id="s2_1_3">
<label>2.1.3</label>
<title>Anthocyanins</title>
<p>Anthocyanins, such as cyanidin, delphinidin, malvidin, pelargonidin, and peonidin, are pigments commonly found in leaves, flowers, and fruits resulting from the hydroxylation, methylation, glycosylation or acylation of the C-ring of the basic skeleton of flavonoids (<xref ref-type="bibr" rid="B6">Alappat and Alappat, 2020</xref>). They can be found in outer cell layers of fruits, like berries and red grapes (<xref ref-type="bibr" rid="B165">Panche et&#xa0;al., 2016</xref>), and are predominantly found in sprouts and flowers of buckwheat, being considered as the predominant group of polyphenols in pigmentation (<xref ref-type="bibr" rid="B65">Fang et&#xa0;al., 2019</xref>). Their functions in plants, as buckwheat, include attracting pollinators (<xref ref-type="bibr" rid="B112">Khan et&#xa0;al., 2022a</xref>), UV protection (<xref ref-type="bibr" rid="B42">Costa et&#xa0;al., 2015</xref>), antioxidant activity (<xref ref-type="bibr" rid="B222">Tsurunaga et&#xa0;al., 2013</xref>), and stress response (<xref ref-type="bibr" rid="B135">Li et&#xa0;al., 2015</xref>).</p>
</sec>
<sec id="s2_1_4">
<label>2.1.4</label>
<title>Flavonols</title>
<p>Flavonols are compounds glycosylated in the C-3 position of the C-ring of the basic skeleton (<xref ref-type="bibr" rid="B69">Flamini et&#xa0;al., 2013</xref>). The most known flavonols in buckwheat are rutin, quercetin and kaempferol (<xref ref-type="bibr" rid="B21">Barreca et&#xa0;al., 2021</xref>). They behave as UV- and photo-protectors, as strongly absorb UV-A and UV-B wavelength (<xref ref-type="bibr" rid="B197">&#x160;ikuten et&#xa0;al., 2020</xref>). They also participate in the regulation of auxin transport and cellular redox status, being involved in physiological processes as root development and gravitropism (<xref ref-type="bibr" rid="B78">Gayomba et&#xa0;al., 2016</xref>), shoot development (<xref ref-type="bibr" rid="B123">Kuhn et&#xa0;al., 2016</xref>), stomatal movements (<xref ref-type="bibr" rid="B236">Watkins et&#xa0;al., 2017</xref>), and, ultimately, in plant growth. Flavonols also play an important role in antioxidant activities by inhibiting ROS synthesis or by scavenging ROS after oxidative burst (<xref ref-type="bibr" rid="B154">Muhlemann et&#xa0;al., 2018</xref>). These metabolites, deeper explained in the section 3 of this review, are the most predominant in buckwheat, and are the main responsible of its pharmaceutical potential.</p>
</sec>
<sec id="s2_1_5">
<label>2.1.5</label>
<title>Flavanones</title>
<p>Flavanones, also present in buckwheat, are generally present in all citrus fruits such as oranges, lemons and grapes, and are responsible of the bitter taste of juice and peel of citrus fruit (<xref ref-type="bibr" rid="B246">Zhao et&#xa0;al., 2020</xref>). Hesperitin, naringenin and eriodyctiol are examples of this class of flavonoids (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). In particular, naringenin is a very special molecule in buckwheat, not only because plays a key role as precursor of other flavonoids, such as quercetin, rutin, kaempferol etc (<xref ref-type="bibr" rid="B143">Mar&#xed;n et&#xa0;al., 2018</xref>), but also because these flavanones confer potential human health benefits such us antioxidant, anticancer, anti-inflammatory, blood lipid-lowering and cholesterol-lowering (<xref ref-type="bibr" rid="B165">Panche et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B207">Stabrauskiene et&#xa0;al., 2022</xref>). In fact, it has been recently characterized as a promising treatment strategy against COVID&#x2010;19 due to its antiviral and anti&#x2010;inflammatory effects (<xref ref-type="bibr" rid="B223">Tutunchi et&#xa0;al., 2020</xref>). Regarding the importance in plants, flavanones play essential roles in insect-plant interactions, pigmentation, heavy metal tolerance, disease resistance, and UV-protection (<xref ref-type="bibr" rid="B113">Khan and Dangles, 2014</xref>).</p>
</sec>
<sec id="s2_1_6">
<label>2.1.6</label>
<title>Flavanols</title>
<p>Flavanols are a subclass of flavonoids with a hydroxyl group on either C-3 or C-4. There are four main types: flavan-3-ols, flavan-4-ols, isoflavan-4-ols, and flavan-3,4-ols. The most common flavanols in buckwheat are catechin and epicatechin, followed by procyanidin (<xref ref-type="bibr" rid="B102">Janovsk&#xe1; et&#xa0;al., 2021</xref>). Regarding human health, flavanols have been shown to improve vascular function through the enhance of endothelial function, resulting in the improvement of blood flow and the reduction of inflammation in the blood vessels (<xref ref-type="bibr" rid="B7">Al-Dashti et&#xa0;al., 2018</xref>), and in cognitive functions (<xref ref-type="bibr" rid="B164">Pa and Gazzaley, 2014</xref>). In plants, flavanols play an important role in defense against pathogens and herbivores through antimicrobial activity properties (<xref ref-type="bibr" rid="B147">Medic et&#xa0;al., 2021</xref>), flower pigmentation, UV protection by absorbing harmful UV radiation, and as ROS scavengers avoiding DNA damage and oxidative stress (<xref ref-type="bibr" rid="B43">Csepregi and Hideg, 2018</xref>). Flavanol content in leaves is inversely proportional to anthocyanins content, and flowers with high flavanol contents are mainly cream-white (<xref ref-type="bibr" rid="B157">Nakayama, 2020</xref>; <xref ref-type="bibr" rid="B89">Han et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s2_1_7">
<label>2.1.7</label>
<title>Flavones</title>
<p>Flavones are characterized by their chemical structure, which includes a backbone of 2-phenylchromen-4-one. The name &#x201c;flavone&#x201d; is derived from the Latin word &#x201c;<italic>flavus</italic>&#x201d;, referring to the yellow pigmentation often associated to these compounds (<xref ref-type="bibr" rid="B51">Deryabin et&#xa0;al., 2019</xref>). Flavones are widely present in leaves, flowers and fruits as glucosides. Apigenin, luteolin, vitexin and orientin are the most common flavones in buckwheat (<xref ref-type="bibr" rid="B102">Janovsk&#xe1; et&#xa0;al., 2021</xref>). Like other flavonoids, flavones serve to a variety of purposes that help plants to adapt to different biological environments, such as (i) defense against UV radiation and oxidative stress; (ii) interspecies interactions (pathogen resistance, symbiosis, defense against herbivory, and allelopathy); and (iii) plant development (co-pigmentation with anthocyanins, and lignification); among others (<xref ref-type="bibr" rid="B103">Jiang et&#xa0;al., 2016</xref>).</p>
</sec>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Other phenolic compounds</title>
<p>Phenolic acids, stilbenes, lignans, tannins, and curcuminoids are phenolic compounds structurally different from flavonoids. They are characterized by the absence of the flavonoid structure, and typically contain a single phenol ring or other unique structures. While most of the phenolic compounds in this group have smaller and simpler chemical structures than flavonoids, there are some phenolic compounds with complex structure and high molecular weight, as well. Phenolic acids have one carboxylic group and one or more OH groups instead of the single phenyl group (<xref ref-type="bibr" rid="B221">Tsimogiannis and Oreopoulou, 2019</xref>). Phenolic acids can be further classified into two main groups: hydroxycinnamic acids (derived from cinnamic acid) and hydroxybenzoic acids (derived from benzoic acid). Examples of hydroxycinnamic acids include ferulic, caffeic, and <italic>p</italic>-coumaric acids, while examples of hydroxybenzoic acids include gallic and protocatechuic acids, both very ubiquitous in buckwheat (<xref ref-type="bibr" rid="B121">Koval et&#xa0;al., 2020</xref>), contributing to the color, flavor, and nutritional properties of this pseudo-cereal. Moreover, phenolic acids are known for their antioxidant activity. They can scavenge free radicals and other reactive oxygen species, helping to protect cells from oxidative damage. Phenolic acids have been also associated with various health benefits, as anti-inflammatory, antimicrobial, and anticancer properties (<xref ref-type="bibr" rid="B77">Garc&#xed;a-P&#xe9;rez et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B61">Embuscado, 2015</xref>; <xref ref-type="bibr" rid="B85">Gon&#xe7;alves et&#xa0;al., 2017</xref>).</p>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>Most abundant phenolic compounds in buckwheat</title>
<p>Nowadays, buckwheat is mainly cultivated for its versatile and highly nutritious seeds, which are used for gluten free bread, pasta, tea, flour, etc. However, the cultivation and study of seedlings and adult plants (sprouts, shoots and roots) have gained importance in the recent years due to their high content of phenolic compounds (PCs), which are very beneficial for plant, animal and human health (<xref ref-type="bibr" rid="B110">Karki et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B60">Dziadek et&#xa0;al., 2018</xref>). The most abundant PCs found in the two major cultivated species of buckwheat, <italic>F. esculentum</italic> and <italic>F. tataricum</italic>, are flavonoids and phenolic acids (<xref ref-type="bibr" rid="B141">Mansur et&#xa0;al., 2022</xref>).</p>
<sec id="s3_1">
<label>3.1</label>
<title>Rutin</title>
<p>The most abundant PC in buckwheat is the flavonol rutin, found in both species, but specially in <italic>F. tataricum</italic> (<xref ref-type="bibr" rid="B150">Mikulajov&#xe1; et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B228">Vieites-&#xc1;lvarez et&#xa0;al., 2023a</xref>). Rutin is a glycoside of the flavonoid quercetin, as is a combination of this flavonol with the hydroxyl group at position C-3 substituted with glucose and rhamnose sugar groups (disaccharide rutinose) (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). <xref ref-type="bibr" rid="B171">Polekhina (2013)</xref> revealed that rutin is mainly found in the leaves and flowers of buckwheat plants (between 63.2 to 76.5% of the total amount of rutin in a plant) followed by the stems (from 7 to 15.7%), and finally the roots (from 1.7 to 5%), depending on the variety. <xref ref-type="bibr" rid="B29">Borovaya and Klykov (2020)</xref> found a similar pattern of accumulation, with large amounts of rutin in flowers of <italic>F. esculentum</italic> (47 to 63 mg g<sup>-1</sup>), significantly less in stems (6 to 14 mg g<sup>-1</sup>), and the minimal amount in roots (3 to 8 mg g<sup>-1</sup>). Moreover, <xref ref-type="bibr" rid="B108">Kalinov&#xe1; et&#xa0;al. (2019)</xref> also found great amounts of rutin in the embryo axis with the cotyledons.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Chemical structure of rutin (quercetin + disaccharide rutinose).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1354672-g004.tif"/>
</fig>
<p>Rutin has shown great benefits for human health such as antioxidant, antihypertensive and anti-inflammatory compound (<xref ref-type="bibr" rid="B162">Nishimura et&#xa0;al., 2016</xref>). Into the plant, rutin is involved in the reduction of environmental stress, UV-light protection, antioxidant activity and disease resistance (<xref ref-type="bibr" rid="B212">Suzuki et&#xa0;al., 2015</xref>). <xref ref-type="bibr" rid="B231">Vollmannov&#xe1; et&#xa0;al. (2013)</xref> studied the amount of rutin, and its antioxidant activity, in the pseudo-cereals buckwheat, quinoa and amaranth, and showed that the large amount of rutin found in buckwheat is probably correlated to the strong antioxidant activity of this crop, which is higher than quinoa or amaranth. In the same way, <xref ref-type="bibr" rid="B132">Lee et&#xa0;al. (2016)</xref> found that rutin and orientin contents in buckwheat are directly correlated to its antioxidant properties, which are closely linked to the attenuation of the impact of environmental stress on plant metabolism. <xref ref-type="bibr" rid="B203">Song et&#xa0;al. (2022)</xref> tested two <italic>F. tataricum</italic> varieties (named TR and CG) at different altitudes, and found that the synthesis of rutin in TR was upregulated as response to cold stress. Moreover, rutin is also considered a strong phytotoxic molecule. <xref ref-type="bibr" rid="B128">Ladhari et&#xa0;al. (2020)</xref> showed that high levels of rutin inhibited the growth of <italic>Lactuca sativa</italic> (L.), <italic>Raphanus sativus</italic> (L.), <italic>Peganum harmala</italic> (L.) and <italic>Silybum marianum</italic> (L.). The exposure of monocotyledonous and dicotyledonous plants such as alfalfa, cress, lettuce, rapeseed, barnyard grass, foxtail fescue, Italian ryegrass, and timothy to aqueous extracts of <italic>Cassia alata</italic> (L.), a plant species with high rutin content, resulted in the growth inhibition of shoots and roots for all the species (<xref ref-type="bibr" rid="B47">Das et&#xa0;al., 2019</xref>). Additionally, rutin, together with chlorogenic acid, were responsible for the inhibition of the growth of <italic>Allium cepa</italic> L (<xref ref-type="bibr" rid="B72">Fonseca et&#xa0;al., 2017</xref>).</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Quercetin</title>
<p>Quercetin, the precursor of rutin, is other important PC in buckwheat. Quercetin is a type of flavonol with a hydroxyl group on the third carbon of the C-ring (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>) that is mainly found in seeds and cotyledons of buckwheat, especially in Tartary buckwheat. <xref ref-type="bibr" rid="B211">Suzuki et&#xa0;al. (2005)</xref> found a negative correlation between rutin and quercetin contents, as quercetin is an intermediate compound in the rutin biosynthesis in plants. In this sense, <xref ref-type="bibr" rid="B178">Ren and Sun (2014)</xref> observed that quercetin content started to decrease five days after sprouting, when rutin content started to increase. Quercetin participates in numerous physiological processes in plants, including photosynthesis, pollen development, antioxidant machinery, and seed germination (<xref ref-type="bibr" rid="B198">Singh et&#xa0;al., 2021</xref>). <xref ref-type="bibr" rid="B127">Kurepa et&#xa0;al. (2016)</xref> discovered that <italic>A. thaliana</italic> treated with quercetin reduced the amount of ROS produced by the ROS-producing herbicide paraquat, suggesting that quercetin would be acting as an antioxidant. <xref ref-type="bibr" rid="B159">Ng et&#xa0;al. (2015)</xref> and <xref ref-type="bibr" rid="B217">Tohge et&#xa0;al. (2016)</xref> also attributed to quercetin the ability to promote healthy plant growth and development by controlling auxin transport, limiting cell proliferation, and promoting the cell elongation phase. Due to its potent antioxidant properties, quercetin effectively protects plants from a variety of biotic and abiotic stress factors (<xref ref-type="bibr" rid="B198">Singh et&#xa0;al., 2021</xref>). Quercetin has also shown potential to interfere with the development of other species. This flavonoid induced the inhibition of germination of alfalfa seeds and the reduction in the weight and length of alfalfa seedlings (<xref ref-type="bibr" rid="B82">Ghimire et&#xa0;al., 2019</xref>). Moreover, quercetin is very effective inhibiting root development. <xref ref-type="bibr" rid="B67">Fern&#xe1;ndez-Aparicio et&#xa0;al. (2021)</xref> found that root growth of <italic>Phelipanche ramose</italic> (L.) Pomel was inhibited by <italic>F. esculentum</italic> root exudates with high content of quercetin. <xref ref-type="bibr" rid="B40">Coelho et&#xa0;al. (2017)</xref> observed that quercetin extracts had the same effects in primary roots of leguminous plants. Additionally, <xref ref-type="bibr" rid="B35">Chen et&#xa0;al. (2019)</xref> and <xref ref-type="bibr" rid="B245">Zhao et&#xa0;al. (2023)</xref> discovered that quercetin has potential to affect photosynthesis, respiration, cell membrane, and enzymatic system of <italic>Microcystis aeruginosa</italic> Kutzing&#x2019;s, causing oxidative damage in a concentration-dependent way.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Quercetin chemical structure.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1354672-g005.tif"/>
</fig>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Phenolic acids</title>
<p>Phenolic acids, including chlorogenic, caffeic, vanillic, ferulic, <italic>p</italic>-coumaric, gallic, dihydroxybenzoic, syringic, protocatechuic, salicylic, phthalic and <italic>m</italic>-toluic acids, among others, are also very important compounds in buckwheat tissues (<xref ref-type="bibr" rid="B232">Vollmannov&#xe1; et&#xa0;al., 2021</xref>). The function of phenolic acids in plants is very diverse, being involved in various aspects of plant growth, development, and defense. Phenolic acids serve as signaling molecules; especially salicylic acid (<xref ref-type="bibr" rid="B117">Kobyletska et&#xa0;al., 2022</xref>), which plays a role in plant-microbe interactions (<xref ref-type="bibr" rid="B140">Mandal et&#xa0;al., 2010</xref>) and contribute to plant resistance responses to stress (<xref ref-type="bibr" rid="B213">Sytar et&#xa0;al., 2014</xref>). In general, phenolic acids show antioxidant properties acting as protective agents against oxidative stress (<xref ref-type="bibr" rid="B239">Wiczkowski et&#xa0;al., 2016</xref>). Additionally, these specialized compounds are involved in regulating enzymatic activity, influencing growth and development processes, attracting pollinators and seed dispersers, and providing UV protection (<xref ref-type="bibr" rid="B37">Cheynier et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B142">Marchiosi et&#xa0;al., 2020</xref>). Therefore, these versatile compounds are essential for plant survival and adaptation to their environment. Most of them has shown potential to manage surrounding weeds. According to <xref ref-type="bibr" rid="B92">Hoang-Anh et&#xa0;al. (2021)</xref>, the mode of action of phenolic acids could involve the induction of oxidative stress in target plants, the modification of cell division and permeability, the alteration of photosynthesis, respiration, and transpiration, and the modulation of gene expression, protein biosynthesis, phytohormone activities, and enzyme functions. For example, <xref ref-type="bibr" rid="B73">Fu et&#xa0;al. (2019)</xref> observed allelopathic effects of phenolic acids on seedling growth and photosynthesis of <italic>Rhododendron delavayi</italic> Franch, and found that ferulic, chlorogenic and protocatechuic acids were highly effective inhibiting stomatal activity (opening, conductance, density and length and width), and reducing the photosynthetic rate. As well, they demonstrated that chlorogenic acid significantly inhibited conductance and transpiration rate in <italic>R. delavayi</italic> seedlings. Leaf extracts of <italic>Juglans regia</italic> containing, among other compounds, protocatechuic and caffeic acids, were able to inhibit <italic>Amaranthus retroflexus</italic> L. and <italic>C. album</italic> L. germination and seedling growth (<xref ref-type="bibr" rid="B55">&#x110;or&#x111;evi&#x107; et&#xa0;al., 2022</xref>). Moreover, leaf litter leachates of different alfalfa cultivars with high content of <italic>p</italic>-hydroxybenzoic, caffeic, <italic>p</italic>-coumaric, chlorogenic, ferulic and cinnamic acids, inhibited seedling growth of <italic>Festuca arundinacea</italic> (Schreb.) Darbysh, and <italic>Sorghum sudanense</italic> (Piper) Stapf (<xref ref-type="bibr" rid="B235">Wang et&#xa0;al., 2017</xref>).</p>
</sec>
</sec>
<sec id="s4">
<label>4</label>
<title>Using buckwheat in agroecology and organic farming for weed management</title>
<p>The fast establishment of buckwheat in the environment, its high canopy growth, and the presence of phenolic compounds with antioxidant but also allelopathic properties, makes of buckwheat a very interesting crop to be exploited on sustainable agricultural such us organic farming and agroecology (<xref ref-type="bibr" rid="B208">Sturm and Gerhards, 2016</xref>). These types of farming are focused on the production of food without the use of synthetic fertilizers, pesticides, other artificial chemicals or genetically modified organisms (GMOs) (<xref ref-type="bibr" rid="B99">International Federation of Organic Agriculture Movements IFOAM, 2020</xref>). Organic farming is linked to several benefits to environment and human health (<xref ref-type="bibr" rid="B58">Dur&#xe1;n-Lara et&#xa0;al., 2020</xref>), being an environmentally sustainable solution to prioritize the protection of soil health, water quality, and biodiversity (<xref ref-type="bibr" rid="B196">Siddique et&#xa0;al., 2014</xref>). Beyond the same purpose, agroecology entails the application of the science of ecology to the study, design, and management of sustainable food systems, the integration of the diverse knowledge systems generated by food system professionals, and the involvement of the social movements that are promoting the transition to fair, just, and responsible food systems (<xref ref-type="bibr" rid="B15">Anderson et&#xa0;al., 2019</xref>).</p>
<p>In this context, the buckwheat&#x2019;s inherent properties make it an ideal crop for organic farming and agroecology, as its growth and specialized metabolites favor soil health and quality while sustainably managing weeds. On the one hand, due to its deep and fibrous root system, buckwheat roots help to improve soil structure, increasing porosity and allowing better infiltration of water and air (<xref ref-type="bibr" rid="B96">Hudek et&#xa0;al., 2021</xref>). On the other hand, the fast shoots development helps to avoid wind and water erosion by providing a surface cover and improving water infiltration. Moreover, this property is also related to the addition of organic matter to the soil through decomposition of its biomass (<xref ref-type="bibr" rid="B153">Mpanga, 2022</xref>). Its short flowering period promotes increased biodiversity as buckwheat flowers are able to attract a variety of pollinators, such as bees and butterflies, which contribute to the pollination of other nearby crops and can provide shelter and food for beneficial insects, such as pest predators, helps to maintain a natural balance in the agricultural ecosystem (<xref ref-type="bibr" rid="B169">P&#x142;a&#x17c;ek et&#xa0;al., 2023</xref>). Finally, buckwheat can help in weed control in organic farming. Due to its rapid growth, dense cover and high presence of specialized compounds in its tissues, which can behave as allelochemical compounds through root exudates, leaf leachates or litter decomposition, buckwheat can compete with weeds, reducing their growth and spread (<xref ref-type="bibr" rid="B63">Falquet et&#xa0;al., 2015</xref>).</p>
<sec id="s4_1">
<label>4.1</label>
<title>Allelopathic potential of buckwheat for weed management</title>
<p>Allelopathy, defined by <xref ref-type="bibr" rid="B180">Rice (1984)</xref> in 1984, refers to both inhibitory and stimulatory reciprocal biochemical interactions between plants and microorganisms including any direct or indirect effect (by microorganisms) of one plant on another through the production of chemical compounds. Allelopathy plays an important role in agroecosystems through a wide variety of interactions between plants and other organisms (<xref ref-type="bibr" rid="B17">Aslam et&#xa0;al., 2017</xref>).</p>
<p>The screening of allelopathic buckwheat cultivars has been of great importance during the past decade. Allelopathic crops naturally exude specialized metabolites (phenolic acids and flavonoids) that can be useful for sustainably manage surrounding weeds, helping to reduce the use of synthetic herbicides by farmers (<xref ref-type="bibr" rid="B17">Aslam et&#xa0;al., 2017</xref>).</p>
<p>The allelopathic effects of buckwheat are attributed to the compounds present in the plant, including flavonoids, phenolic acids, fatty acids, and other allelochemicals (<xref ref-type="bibr" rid="B137">Li et&#xa0;al., 2019</xref>). The release of these compounds to the medium can affect the growth and development of surrounding weeds (<xref ref-type="bibr" rid="B228">Vieites-&#xc1;lvarez et&#xa0;al., 2023a</xref>, <xref ref-type="bibr" rid="B229">Vieites-&#xc1;lvarez et&#xa0;al., 2023b</xref>).</p>
<p>Buckwheat plants start to release allelopathic compounds during germination and early development (<xref ref-type="bibr" rid="B107">Kalinov&#xe1; et&#xa0;al., 2005</xref>). <xref ref-type="bibr" rid="B249">Zou et&#xa0;al. (2014)</xref> also found that allelochemicals released by buckwheat in an early phase of development, induced the growth inhibition of couch grass (<italic>Elymus repens</italic> L.), showing the weed-suppressive ability of buckwheat trough root exudates. As well, <xref ref-type="bibr" rid="B81">Gfeller et&#xa0;al. (2018)</xref> showed that common buckwheat could alter its root exudation after recognizing the presence of <italic>A. retroflexus</italic>, suppressing this weed&#x2019; growth. <xref ref-type="bibr" rid="B67">Fern&#xe1;ndez-Aparicio et&#xa0;al. (2021)</xref> indicated that buckwheat root exudates, rich in the flavonoid quercetin and the stilbene <italic>p-</italic>coumaric, were able to induce the inhibition in <italic>Phelipanche ramosa</italic> (L.) Pomel radicles. More recently, <xref ref-type="bibr" rid="B229">Vieites-&#xc1;lvarez et&#xa0;al. (2023b)</xref> tested the allelopathic potential of four buckwheat varieties (Kora, Eva, PI481671 and Gema) from two different species (<italic>F. esculentum</italic> and <italic>F. tataricum</italic>), and found that buckwheat plants growing in co-culture with the weeds <italic>L. rigidum</italic> or <italic>P. oleracea</italic> released to the medium much higher amounts of flavonoids (i.e., rutin, quercetin, orientin, vitexin, etc.) and phenolic acids (i.e., ferulic acid, protocatechuic acid, syringic acid, etc.) during the first days of development (7-17 days), than buckwheat plants growing alone, which favored the inhibition of weeds&#x2019; growth. These allelochemicals present in different tissues are often extracted to test their inhibitory potential on weeds.</p>
<p>
<xref ref-type="bibr" rid="B151">Mioduszewska et&#xa0;al. (2013)</xref> evaluated water extracts from common buckwheat (from aboveground and roots tissues), and found that aboveground extracts showed stronger inhibition in the germination of lettuce seeds than root extracts. <xref ref-type="bibr" rid="B224">Uddin et&#xa0;al. (2013)</xref> observed that 7.5 mg mL<sup>-1</sup> Tartary buckwheat extract reduced the growth of <italic>Galium spurium</italic> L. up to 33.7%. Recently, <xref ref-type="bibr" rid="B214">Szwed et&#xa0;al. (2020)</xref> showed that a very low percentage (1%) of the aqueous extract of 14-day common buckwheat plants was able to inhibit root development of wild oat (<italic>Avena fatua</italic> L.), yellow foxtail (<italic>Setaria glauca</italic> L. P. Beauv.), barnyardgrass (<italic>Echinochloa crus-galli</italic> (L.) Beauv.), common windgrass (<italic>Apera spica-venti</italic> (L.) P. Beauv.), catchweed bedstraw (<italic>Galium aparine</italic> L.), scentless mayweed (<italic>Matricaria inodora</italic> L.) and gallant soldier (<italic>Galinsoga parviflora</italic> Cav.), and the shoot length of wild oat. As well, more recently (<xref ref-type="bibr" rid="B33">Cechin et&#xa0;al., 2023</xref>), three concentrations of extracts from seeds or shoots of buckwheat (25, 50 and 100%) resulted in the reduction of germination and abnormal seedling development of the weeds <italic>B. pilosa</italic> and <italic>Euphorbia heterophylla</italic> L., with <italic>B. pilosa</italic> being more sensitive than <italic>E. heterophylla</italic> to both extracts of buckwheat.</p>
<p>This phytotoxic activity of buckwheat is probably due to the PCs present in buckwheat tissues, which can be accumulated in shoots or roots or relocated to be released to the medium (<xref ref-type="bibr" rid="B230">Vieites-&#xc1;lvarez et&#xa0;al., 2023c</xref>). In other research, gallic acid and (+)-catechin, extracted from buckwheat shoots, were found to be the responsible for the inhibition of roots and shoots of lettuce seedlings (<xref ref-type="bibr" rid="B100">Iqbal et&#xa0;al., 2017</xref>). A recent study demonstrated that methanolic extracts of gallic, caffeic and ferulic acids in concentrations of 153.96, 69.13, and 39.80 ppm, respectively, were enough for inhibiting the growth of <italic>Chenopodium murale</italic> L. and <italic>A. viridis</italic> L (<xref ref-type="bibr" rid="B12">Alsharekh et&#xa0;al., 2022</xref>). <xref ref-type="bibr" rid="B191">&#x160;&#x107;epanovi&#x107; et&#xa0;al. (2022)</xref> evaluated the phytotoxic potential of several PCs (chlorogenic, caffeic, ferulic, gallic, protocatechuic, <italic>p</italic>-hydroxybenzoic, syringic, vanillic, and <italic>p</italic>-coumaric acids) on <italic>Ambrosia artemisiifolia</italic> L. and found that treatment with ferulic acid, vanillic acid, <italic>p</italic>-coumaric acid, <italic>p</italic>-hydroxybenzoic acid, or a mixture of all phenolic acids, highly inhibited early growth parameters, suggesting the strong phytotoxic potential of these compounds and the higher toxicity of phenolic acids mixtures. The research of <xref ref-type="bibr" rid="B81">Gfeller et&#xa0;al. (2018)</xref> proposed that these compounds would be the responsible for the suppression caused by common buckwheat to pigweed, goosefoot and barnyard grass growth by 53, 42 and 77%, respectively, as they observed significant growth inhibition in an experiment without physical root interactions.</p>
<p>In this context, the screening and selection of varieties with strong allelopathic potential, both through root exudates of living plants or through metabolites present in plant tissues, could provide an alternative solution to the use of synthetic herbicides, as it has been shown that buckwheat has the ability to sustainably manage competing surrounding weeds. Furthermore, the proven effectiveness of these specialized metabolites makes buckwheat a very suitable crop for other agroecological cultivation methods such as mulching, cover cropping, intercropping and green manure.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Competition role in weed management</title>
<p>Although, nowadays, there are numerous studies on the potential of allelopathy or allelochemicals to manage weeds, it is very difficult, under natural conditions, to differentiate between allelopathy and competition, as both occur simultaneously in plant communities and co-exist in the agroecosystems influencing weed germination and development. Laboratory tests usually do not take into consideration the soil environment and the multiple abiotic conditions that crops and weeds cope with during their development in the agroecosystems. Moreover, specialized metabolites undergo continuous changes under natural conditions depending on the physical, chemical and biological conditions of the soil, which will determine their phytotoxic level. Therefore, the interaction of weeds and crops in communities is a complex combination of competition for resources and the effect of plant specialized metabolites (<xref ref-type="bibr" rid="B20">B&#x105;czek and Halarewicz, 2022</xref>).</p>
<p>Regarding resource competition, weeds can compete with crops for water, nitrogen, light, nutrients and space (<xref ref-type="bibr" rid="B186">Sardana et&#xa0;al., 2017</xref>). The establishment of plants in the environment depends heavily on phenotypic traits like shoot and root length, leaf number, and plant height (<xref ref-type="bibr" rid="B125">Kunstler et&#xa0;al., 2016</xref>). Crop accessions with a higher initial growth rate and a quicker canopy development rate can establish more quickly and will be less impacted by weed competition (<xref ref-type="bibr" rid="B49">Dass et&#xa0;al., 2017</xref>). <xref ref-type="bibr" rid="B210">Sturm et&#xa0;al. (2018)</xref> evaluated the relative proportion of allelopathic effects with respect to weed inhibition. They used <italic>F. esculentum</italic> as a cover crop with and without activated charcoal in the soil (used as adsorbent of allelopathic substances), and concluded that although allelopathy contributed significantly to the overall suppression of weeds, competition had a bigger impact. These authors concluded that to increase the effectiveness of weed control, an allelopathic cover crop should show competitive prerequisites, as rapid germination, fast development, dense canopy, and high soil coverage.</p>
<p>Common buckwheat is one of the fastest growing cover crops used in Western Europe. Seedlings emerge in less than one&#x2009;week, and cover 40&#x2013;80% of the soil within 4&#x2009;weeks after sowing, producing around 200&#x2009;g&#x2009;m<sup>&#x2212;2</sup> of shoot and root dry matter. For this reason, buckwheat is considered one of the best cover crops (<xref ref-type="bibr" rid="B31">Brust et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B126">Kunz et&#xa0;al., 2016</xref>).</p>
<p>During the past decades, some authors suggested that light competition could be the most important factor for weed suppression in buckwheat fields, as the rapid development of buckwheat canopy prevents weed establishment due to lack of light (<xref ref-type="bibr" rid="B219">Tominaga and Uezu, 1995</xref>; <xref ref-type="bibr" rid="B25">Bicksler and Masiunas, 2009</xref>). However, there has been no experimental evidence for this hypothesis so far. In 2013, <xref ref-type="bibr" rid="B79">Gebhard et&#xa0;al. (2013)</xref>, demonstrated that the amount of crop biomass produced (i.e., the amount of shading), was closely correlated to the ability of legumes to compete with weeds. <xref ref-type="bibr" rid="B31">Brust et&#xa0;al. (2014)</xref> conducted a field experiment with cover crops such as white mustard, oilseed radish, forage radish, phacelia, Tartary buckwheat, red oat, and grain amaranth, and a no cover crop control to identify the weed suppression ability of cover crops. They found that Tartary buckwheat was the cover crop with the highest shoot growth eight weeks after sowing, resulting in more than 95% weed suppression. <xref ref-type="bibr" rid="B249">Zou et&#xa0;al. (2014)</xref> concluded that the high leaf area index (LAI) of buckwheat could be one of the weed-suppressive factors decreasing <italic>E. repens</italic> growth and lowering its weight dry mass. As well, <xref ref-type="bibr" rid="B199">Smith et&#xa0;al. (2020)</xref> obtained a 98% weed suppression in a buckwheat monoculture field due to its cover soil ability.</p>
<p>Moreover, competition not only takes place aboveground. Below-ground competition for resources constitutes also an important aspect of crop-weed interaction. This type of competition is naturally occurring for space, soil nutrients and water, as plants mainly take up nutrients and water from the proximal root zone (<xref ref-type="bibr" rid="B16">Asaduzzaman et&#xa0;al., 2014</xref>). <xref ref-type="bibr" rid="B64">Falquet et&#xa0;al. (2014)</xref> performed an experiment in pot trials in order to detect competition in soil between two varieties of buckwheat and redroot pigweed. They found that the roots of redroot pigweed were up to 87% smaller after growing in co-culture with the buckwheat varieties than after growing alone in the same space (no influence of space limitation). Recently, <xref ref-type="bibr" rid="B228">Vieites-&#xc1;lvarez et&#xa0;al. (2023a)</xref> showed that some buckwheat varieties could modify their growth resulting in longer roots after co-cultivation with monocot or dicot weeds. The ability to modify root length could be an advantage in water uptake competition. In this sense, <xref ref-type="bibr" rid="B144">Mart&#xed;nez-Go&#xf1;i et&#xa0;al. (2023)</xref> tested the behavior of wheat (<italic>Triticum aestivum</italic> L.), spelt (<italic>Triticum spelta</italic> L.), and common buckwheat (<italic>F. esculentum</italic>) under well-watered conditions, mild drought an extreme drought, and found that buckwheat was the only crop able to cope with extreme drought conditions by increasing its water use efficiency and maintaining its photosynthetic parameters. This property of buckwheat is, and in the future will be even more, an advantage in increasingly common and widespread periods of drought due to climate change (<xref ref-type="bibr" rid="B242">Yuan et&#xa0;al., 2023</xref>).</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Allelopathy and competition interaction in weed management strategies (cover crops, mulching, intercropping, crop rotation and green manure)</title>
<p>The combination of allelopathic and competitive traits in buckwheat as a cover crop may enhance its weed suppressive capacity (<xref ref-type="bibr" rid="B71">Florence et&#xa0;al., 2019</xref>), making this pseudo-cereal suitable for other weed management methods such as intercropping, crop rotation or green manuring. These techniques are often indicated as distinct and separate methods, but they can be considered as sister strategies, as all are based in the same buckwheat properties (<xref ref-type="bibr" rid="B189">Scavo and Mauromicale, 2021</xref>). In fact, cover crops, as buckwheat, can be applied as living mulches, when intercropped with industrial crops; as dead mulches, when plant residues are left on the surface of the soil; or as green manures, when the residues are incorporated into the soil (<xref ref-type="bibr" rid="B189">Scavo and Mauromicale, 2021</xref>). Unfortunately, these techniques have not been further studied in buckwheat during the last decade, although all are promising strategies for the establishment of agroecological approaches in agricultural production, which should be taken into consideration in the near future.</p>
<sec id="s4_3_1">
<label>4.3.1</label>
<title>Mulching</title>
<p>Mulching is the practice of applying a layer of material on top of the soil to provide various benefits to plants and the overall soil. Mulch can be made from a variety of organic or inorganic materials, such as grass clippings, straw, bark chips, stones, plant residues, or plastic. The benefits of mulching include weed suppression, moisture retention, soil temperature control, erosion prevention, nutrient cycling and improved soil structure (<xref ref-type="bibr" rid="B114">Khan et&#xa0;al., 2022b</xref>). Generally, living mulching is a more effective method for controlling weeds than non-living mulching (i.e., cover crop residues), as the first one can suppress weeds throughout the growing season while the second option is more effective in preventing weed seed germination and smothering very young weeds (<xref ref-type="bibr" rid="B237">Westbrook et&#xa0;al., 2022</xref>). Mulches can manage weeds by mixing competition (light, water, and nutrients) and allelopathy. This is a very efficient method for light interception, which compromises germination and growth of many weeds. Moreover, reduction in light intensity is also correlated with modifications in soil temperature, which can also delay the germination of weeds (<xref ref-type="bibr" rid="B22">Batlla and Benech-Arnold, 2014</xref>). Mulches can also modify water availability by transpiration, which could have greater impacts on shallow-rooted weeds than deeper-rooted crops (<xref ref-type="bibr" rid="B237">Westbrook et&#xa0;al., 2022</xref>). Regarding nutrient competition, buckwheat has a high carbon-to-nitrogen (C:N) ratio, which means that it can potentially tie up nitrogen in the soil and temporarily reduce its availability for other plants meanwhile provides a large amount of C to soil (<xref ref-type="bibr" rid="B227">Vecchia et&#xa0;al., 2020</xref>).</p>
<p>Besides, allelopathy plays also an important role in mulching. Buckwheat allelochemicals are present in different plant tissues, as leaves, flowers, shoots and roots, but they are not necessarily released to the medium, accumulating in the different organs of the plant. After the plant cycle, the plant residues can be left in the soil or can be dried to straw. The rapid leaf development of buckwheat allows obtaining more dry matter in less time, making it ideal for this agroecological technique. Root residues from buckwheat were able to inhibit biomass of barnyard grass (<italic>E. crus-galli</italic>) and cleavers (<italic>Galium aparine</italic> L.) in a dose-response way when left on the soil (<xref ref-type="bibr" rid="B215">Szwed et&#xa0;al., 2019</xref>), with different mixtures inducing different weed management potential. The mixture of 0.5&#xa0;t ha<sup>-1</sup> buckwheat with 1.0&#xa0;t ha<sup>-1</sup> marsh pepper residues was the most effective in suppressing weeds while improving grain yields (<xref ref-type="bibr" rid="B3">Afroz et&#xa0;al., 2018</xref>). Mulches of <italic>F. tataricum</italic> tested in a field with <italic>C. album</italic> L., <italic>Matricaria chamomilla</italic> L. and <italic>Stellaria media</italic> (L.) Vill induced a decrease in germination rate and dry matter of those weeds respect to untreated fields (<xref ref-type="bibr" rid="B209">Sturm et&#xa0;al., 2016</xref>). Straw of buckwheat inhibited weed development in broccoli fields too, probably due to the combination of the physical properties of the mulches (color, structure, etc.) and their allelopathic effects (<xref ref-type="bibr" rid="B119">Kosterna, 2014a</xref>). <xref ref-type="bibr" rid="B130">Latify et&#xa0;al. (2017)</xref> proposed that an adequate selection of sunflower competitive varieties in combination with buckwheat living mulch could be the best option to manage weeds on sunflowers fields, as they observed a reduction in the density of weeds (up to 96%) when compared with a control yield (without buckwheat mulches and non-competitive sunflower variety). Moreover, buckwheat helps protecting radish and cucumber crops by suppressing diseases, when added to the soil at least three weeks before planting (<xref ref-type="bibr" rid="B1">Abbasi et&#xa0;al., 2018</xref>). <xref ref-type="bibr" rid="B166">Pfeiffer et&#xa0;al. (2015)</xref>, after a research involving <italic>F. esculentum</italic> mulch and snap beans (<italic>Phaseolus vulgaris</italic> var. Tavera) with bell peppers (<italic>Capsicum annuum</italic> var. Revolution) and fall broccoli (<italic>Brassica oleracea</italic> var. Imperial) as cultivated plants, stated that there is an inverse relation between living mulch biomass and weed biomass, confirming the weed suppression ability of living mulches, although yield productivity could be sometimes reduced due to resource competition among plant species, living mulches and weeds.</p>
<p>Finally, buckwheat mulching not only can be useful for weed control. In conventional cropping systems, buckwheat mulching successfully prevented leaching of herbicides (such as diclosulam and diuron) to the soil by creating a chemical barrier which involves the prevention of soil contamination (<xref ref-type="bibr" rid="B155">Munhoz-Garc&#xed;a et&#xa0;al., 2023</xref>). Moreover, it was observed that after covering the field with buckwheat residues, the daily temperature fluctuations were reduced, and the moisture increased at depths between 0 - 40&#xa0;cm, improving soil broccoli production compared to a field without mulching (<xref ref-type="bibr" rid="B120">Kosterna, 2014b</xref>).</p>
<p>The combination of the two characteristics previously mentioned (presence of phenolic compounds in buckwheat tissues + relevant leaf biomass) makes of buckwheat an ideal crop for mulching.</p>
</sec>
<sec id="s4_3_2">
<label>4.3.2</label>
<title>Intercropping</title>
<p>Intercropping is a strategy defined by the simultaneous growth of two or more crops in the same field over an extended period of time, without necessarily sowing or harvesting them at the same time (<xref ref-type="bibr" rid="B23">Bedoussac et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B136">Li et&#xa0;al., 2020</xref>). Intercropping is an eco-functional practice that has been extensively used to increase crop productivity (<xref ref-type="bibr" rid="B173">Qin et&#xa0;al., 2013</xref>) and land use efficiency (<xref ref-type="bibr" rid="B115">Kidane et&#xa0;al., 2017</xref>), while reducing the quantity of greenhouse gases emitted to the atmosphere when compared with monoculture&#x2019; fields (<xref ref-type="bibr" rid="B158">Naudin et&#xa0;al., 2014</xref>). When allelopathic crops are grown in intercropping systems, allelochemicals are released into the environment through root exudation, volatilization from above or below-ground plant parts, leaching from rainfall, or decomposition of plant debris (<xref ref-type="bibr" rid="B190">Scavo et&#xa0;al., 2018</xref>). By increasing the diversity of the soil&#x2019;s microbial population and facilitating the transport of allelochemicals&#x2019; into the soil, intercropping could improve the allelopathic weed&#x2013;cover crop interactions and, as a result, the phytotoxic effects (<xref ref-type="bibr" rid="B30">Brooker et&#xa0;al., 2015</xref>).</p>
<p>
<xref ref-type="bibr" rid="B26">Biszczak et&#xa0;al. (2020)</xref> found that soybean intercropped with buckwheat (i): 1 row of soybean&#x2009;+&#x2009;1 row of buckwheat, or (ii): 1 row of soybean&#x2009;+&#x2009;2 rows of buckwheat, reached a significant reduction of total weed number and weeds&#x2019; dry weight when compared to mono-cultured soybean fields. Although the highest soybean yields were obtained in intercropping with lentils, it is necessary to choose crops that have a balance between yield improvement and weed control. <xref ref-type="bibr" rid="B36">Cheriere et&#xa0;al. (2020)</xref> found that soybean intercropped with sunflower (2018) and buckwheat (2019) provided the best weed control, while ensuring high soybean yields. <xref ref-type="bibr" rid="B183">Salehi et&#xa0;al. (2018)</xref> intercropped fenugreek and buckwheat in a 2:1 ratio, improving fenugreek productivity. Very recently, <xref ref-type="bibr" rid="B129">Lakshmi et&#xa0;al. (2023)</xref> found that intercropping buckwheat with maize, baby corn, sweet corn and sunflower would work well, and even the microbial activity of the soil may vary depending of the crop mixture.</p>
<p>However, intercropping is not always a good idea. <xref ref-type="bibr" rid="B243">&#x17d;alac et&#xa0;al. (2022)</xref> checked land and water productivity (WP) in an old and dense orchard compared to a younger one, both with intercropping buckwheat or barley and walnut trees, and found a reduction in walnut yields and WP due to competition between crops in old orchards, although buckwheat productivity reached the highest yield and WP in the young orchard. <xref ref-type="bibr" rid="B199">Smith et&#xa0;al. (2020)</xref> compared the efficacy of intercropping six cover crops (with a total of 14 species mixtures) and monoculture these crops to control weed abundance (weed biomass) and to reach weed suppression during three seasons: summer (spring planting-summer termination), fall (summer planting-fall termination), and spring (fall planting-subsequent spring termination). They found that, irrespective of the season, mixtures were never more weed suppressive than the most weed-suppressive cover crop grown as a monoculture. For example, buckwheat monoculture reached a weed suppression of 97-98% in summer season, while the mixture of buckwheat with other crops was less effective ranging from 66 to 96%. Similar patterns were also found for other cover crops, suggesting that farmers could reach better results of weed suppression in monoculture systems.</p>
</sec>
<sec id="s4_3_3">
<label>4.3.3</label>
<title>Crop rotation</title>
<p>Crop rotation is a traditional agricultural practice that offers numerous benefits for weed and pest control, reduced auto-allelopathy and nutrient leaching, improved soil organic matter and soil fertility, and increased crop yields (<xref ref-type="bibr" rid="B189">Scavo and Mauromicale, 2021</xref>). Root exudates and decomposing crop residues from allelopathic buckwheat crops enriched the soil with allelochemicals that favored weed control in the next crop cycle (<xref ref-type="bibr" rid="B101">Jabran et&#xa0;al., 2015</xref>). Buckwheat grows well with other early-maturing crops like potatoes, spring greens, or winter canola. Crop rotation of buckwheat&#x2013;buckwheat&#x2013;potatoes resulted in 30% less N leaching and 16% more tuber yield (<xref ref-type="bibr" rid="B104">Jiang et&#xa0;al., 2022</xref>), maybe due to buckwheat roots containing acyl sucrose, which causes hormonal imbalances in wireworms, whose larvae cause extensive damage to potatoes (<xref ref-type="bibr" rid="B8">Alkhnajari, 2019</xref>).</p>
</sec>
<sec id="s4_3_4">
<label>4.3.4</label>
<title>Green manure</title>
<p>Green manure is a technique that takes advantage of cover crops by harvesting and mixing them with soil to provide valuable organic matter and weed control compounds. Buckwheat has shown to be very useful as green manure due to its allelopathic properties and the ability to incorporate nutrients to the soil, especially phosphorous and nitrogen (<xref ref-type="bibr" rid="B28">Boglaienko et&#xa0;al., 2014</xref>). <xref ref-type="bibr" rid="B145">Masilionyte et&#xa0;al. (2017)</xref> tested the weed-suppressive potential of buckwheat mixed with other cover crop as white mustard (<italic>S. alba</italic>), both cultivated for green manure in a 6-year field experiment. The number of weeds and the weed biomass were significantly lower in buckwheat-mustard mixed fields than after the cultivation of narrow-leafed lupine (<italic>Lupinus angustifolius</italic> L.) in a mixture with oil radish (<italic>Raphanus sativus</italic> L.), thanks to its biomass production and release of allelochemicals into the soil. <xref ref-type="bibr" rid="B177">Reddy et&#xa0;al. (2021)</xref> obtained increased plant growth, dry-matter accumulation, tillers, grain yield, and economic improvement in rice (<italic>Oryza sativa</italic> L.) under green-manuring with buckwheat.</p>
</sec>
</sec>
</sec>
<sec id="s5">
<label>5</label>
<title>Cultivation practices for improving buckwheat potential in weed management</title>
<p>Buckwheat is a crop that can be cultivated under different growing conditions, although different buckwheat species developed different in a diverse range of environments (<xref ref-type="bibr" rid="B176">Rangappa et&#xa0;al., 2023</xref>). Making a selection of the buckwheat cultivars according to the geographical area, weather conditions, need for tillage, or seeding distance could enhance the buckwheat ability to growing better and managing more successfully adventitious plants.</p>
<sec id="s5_1">
<label>5.1</label>
<title>Weather conditions</title>
<p>The temperature, quantity of rainfall and sunshine hours will affect the total biomass production and grain yield and quality of buckwheat (<xref ref-type="bibr" rid="B106">Jung et&#xa0;al., 2015</xref>). Even when buckwheat can increase water use efficiency under extreme drought conditions (<xref ref-type="bibr" rid="B144">Mart&#xed;nez-Go&#xf1;i et&#xa0;al., 2023</xref>), this pseudo-cereal is sensitive to high temperatures and hot dry winds, especially under low humidity conditions (<xref ref-type="bibr" rid="B56">Dra&#x17e;i&#x107; et&#xa0;al., 2016</xref>). However, different species of buckwheat are genetically regulated to better withstand different climatic situations. <italic>F. tataricum</italic> is more resistant to cold due to a DNA methylation (<xref ref-type="bibr" rid="B204">Song et&#xa0;al., 2020</xref>), and more resistant to drought than <italic>F. esculentum</italic>, while this species thrives better on sandy and well-drained soils (<xref ref-type="bibr" rid="B56">Dra&#x17e;i&#x107; et&#xa0;al., 2016</xref>). So, Tartary buckwheat is regarded as a potential crop for cultivation at higher altitudes because of its adaptability to various climatic variables and water-stress regimes, cold temperatures, and nutrient-deficient acid soil (<xref ref-type="bibr" rid="B176">Rangappa et&#xa0;al., 2023</xref>). Temperature and altitude are also one of the most determinant environmental factors in the production of flavonoids as rutin (<xref ref-type="bibr" rid="B200">Sobhani et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B56">Dra&#x17e;i&#x107; et&#xa0;al., 2016</xref>), which will determine the allelopathic properties of buckwheat. Regarding rainfall, buckwheat is highly susceptible to dryness, particularly in early growth stages, during rooting, flowering, and yielding period. However, moisture excess during the later stages of growth has also strong detrimental effects on buckwheat development (<xref ref-type="bibr" rid="B161">Nikolic et&#xa0;al., 2019</xref>).</p>
</sec>
<sec id="s5_2">
<label>5.2</label>
<title>Sowing time</title>
<p>The timing of this activity is quite related to temperature. Sowing time is one non-monetary input that significantly influences crop productivity (<xref ref-type="bibr" rid="B34">Chaudhary et&#xa0;al., 2019</xref>), and buckwheat is not an exception. Sowing time influences seed germination, flowering time, length of vegetation, rutin content, and herb and grain yield of buckwheat (<xref ref-type="bibr" rid="B19">Babu et&#xa0;al., 2018</xref>). The range for seed germination varies from 5 to 42&#xb0;C, but the optimum temperature is around 24-26&#xb0;C (<xref ref-type="bibr" rid="B56">Dra&#x17e;i&#x107; et&#xa0;al., 2016</xref>). Buckwheat seeds must be sown at different times, depending on the geographic area, to avoid flower blasting, which usually occurs at temperatures higher than 32&#xb0;C (<xref ref-type="bibr" rid="B170">P&#x142;a&#x17c;ek et&#xa0;al., 2019</xref>). Once flowering has started, the plants should be at least 10 weeks in non-frost weather conditions. <xref ref-type="bibr" rid="B176">Rangappa et&#xa0;al. (2023)</xref> obtained the highest mean yield of buckwheat when the sowing date was between October and December in the Himalaya region, while in central New York, <xref ref-type="bibr" rid="B27">Bj&#xf6;rkman and Shail (2013)</xref> determined that the optimal timing for sowing buckwheat was late June to early August, and in Korea, <xref ref-type="bibr" rid="B106">Jung et&#xa0;al. (2015)</xref> proposed mid to end of August as the best time for sowing buckwheat.</p>
</sec>
<sec id="s5_3">
<label>5.3</label>
<title>Tillage requirement</title>
<p>To decide on using tillage or a non-till system will mainly depend on the characteristics of the planting soil. Buckwheat, as a low input crop that can be cultivated under reduced tillage system (<xref ref-type="bibr" rid="B39">Chrungoo and Chettry, 2021</xref>). Actually, buckwheat requires low-fertility soil with a moderate nitrogen content to start growing (<xref ref-type="bibr" rid="B75">Gairhe et&#xa0;al., 2015</xref>). Low tillage can boost buckwheat crop germination and establishment by creating a seedbed that facilitates optimal seed-to-soil contact. Additionally, soil&#x2019;s incorporation of organic matter and crop residue can be improved also by tillage, which can enhance the soil&#x2019;s fertility and structure. Tillage can also control the growth of adventitious plant species avoiding the emergence of weeds and burying weed seeds (<xref ref-type="bibr" rid="B156">Myers, 2018</xref>). Nevertheless, buckwheat can be drilled without tillage, which is a viable choice especially for mid-summer planting. This strategy can reduce soil erosion and help preserving soil moisture. Additionally, weed management can also be favored by tillage, because the residue from the previous crop acts as a mulch to prevent the growth of weeds. The particular characteristics of the field, such as the type of the soil, the amount of moisture, and the pressure from weeds, should be taken into account when determining whether or not tillage buckwheat fields (<xref ref-type="bibr" rid="B156">Myers, 2018</xref>).</p>
</sec>
<sec id="s5_4">
<label>5.4</label>
<title>Seed rate, sowing depth and spacing between rows</title>
<p>The optimal seed rate of buckwheat can vary from 90 to 160 grains per m<sup>2</sup> as maximum (<xref ref-type="bibr" rid="B161">Nikolic et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B234">Wan et&#xa0;al., 2023</xref>) because higher seedling rates can result in puny plants with short stems (<xref ref-type="bibr" rid="B218">Tolaini et&#xa0;al., 2016</xref>), which would impede the proper plant development, thus minimizing the ability of buckwheat to control weeds in an agroecological system. A proper plant population is the key for reaching higher crop yields, as plants could use all available resources without inter-cultivar competition (<xref ref-type="bibr" rid="B19">Babu et&#xa0;al., 2018</xref>). Depth of sowing is also crucial for determining the ideal plant population. <xref ref-type="bibr" rid="B80">Gerhards and Schappert (2020)</xref> proposed 2&#x2013;3 cm as a good seeding depth for all cover crop species if a mixture is sown. This allows small and large-seeded cover crops to emerge. <xref ref-type="bibr" rid="B240">Xiang et&#xa0;al. (2014)</xref> tested three different sowing depth in <italic>F. tataricum</italic> (2, 4 and 6&#xa0;cm), and concluded that buckwheat seeds sown at 4&#xa0;cm had an emergence rate, seedling number, seedling rate, stem diameter, dry matter, area per plant, and leaf chlorophyll content significantly higher than when seeds were sown at 2 or 6&#xa0;cm. Regarding row spacing, there are some controversial mainly depending on the species. <xref ref-type="bibr" rid="B118">Kolari&#x107; et&#xa0;al. (2021)</xref> determined that 25&#xa0;cm instead of 50&#xa0;cm spacing between rows resulted in higher average in grain yield for <italic>F. esculentum</italic>. However, <xref ref-type="bibr" rid="B184">Salmankhan et&#xa0;al. (2021)</xref> found that closer spacing of 30&#xa0;cm between rows &#xd7; 10&#xa0;cm between plants resulted in taller plants, while wider row spacing of 45 between rows &#xd7; 15&#xa0;cm between plants resulted in higher number of leaves.</p>
<p>Finally, depending on the intended purpose, the time of harvest may also vary. In a previous study (<xref ref-type="bibr" rid="B109">Kara, 2014</xref>) where buckwheat was harvested at different stages of the crop (1<sup>st</sup> harvest stage (HS): the beginning of flowering; 2<sup>th</sup> HS: 20 days after 1<sup>st</sup> harvest (around the full blooming period); 3<sup>rd</sup> HS: 32 days after 1st harvest (about the end of flowering), 4<sup>th</sup> HS: 45 days after 1<sup>st</sup> harvest (about 25% of grains had turned brown) and 5<sup>th</sup> HS: 55 days after 1<sup>st</sup> harvest (about 50% of grains had turned brown), <xref ref-type="bibr" rid="B109">Kara (2014)</xref> concluded that harvesting time may vary depending on the final interest. For higher plant matter and crop yields (i.e., for mulching, seed production, etc.), buckwheat should be harvested 55 days after sowing, however, for strong mineral nutrient content in plant organs, the plants should be harvested 20 days after sowing (for pharmaceuticals, food sector, etc.).</p>
</sec>
</sec>
<sec id="s6">
<label>6</label>
<title>Buckwheat nutritional profile and health</title>
<p>Buckwheat, in terms of nutrition, has been established as one of the most complete and nutritious foods. Buckwheat grains are a rich source of protein with a balanced amino acid composition, gluten free flour, dietary fiber, vitamins, resistant starch, phytosterols, fagopyrins, fagopyritols, and phenolic compounds (<xref ref-type="bibr" rid="B201">Sofi et&#xa0;al., 2023</xref>). Some slight differences are observed in the nutritional composition of different buckwheat species (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Differences in the nutritional composition of common buckwheat (<italic>Fagopyrum esculentum</italic>) and Tartary buckwheat (<italic>Fagopyrum tataricum</italic>).</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Parameters</th>
<th valign="middle" align="left">Protein</th>
<th valign="middle" align="left">CH</th>
<th valign="middle" align="left">Lipid</th>
<th valign="middle" align="left">Dietary fiber</th>
<th valign="middle" align="left">Ash</th>
<th valign="middle" align="left">Other</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Common buckwheat <xref ref-type="table-fn" rid="fnT1_1">
<sup>a</sup>
</xref>
</td>
<td valign="middle" align="center">12.30</td>
<td valign="middle" align="center">54.50</td>
<td valign="middle" align="center">3.80</td>
<td valign="middle" align="center">7.00</td>
<td valign="middle" align="center">2.00</td>
<td valign="middle" align="center">18.40</td>
</tr>
<tr>
<td valign="top" align="left">Tartary buckwheat <xref ref-type="table-fn" rid="fnT1_1">
<sup>a</sup>
</xref>
</td>
<td valign="middle" align="center">13.15</td>
<td valign="middle" align="center">57.40</td>
<td valign="middle" align="center">3.84</td>
<td valign="middle" align="center">10.6</td>
<td valign="middle" align="center">2.70</td>
<td valign="middle" align="center">10.53</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>CH, carbohydrates; Other refers to other compounds such soluble carbohydrates, organic acid, nucleotides, and other unknown compounds. Data are given in %.</p>
</fn>
<fn id="fnT1_1">
<label>a</label>
<p>Adapted and modified from <xref ref-type="bibr" rid="B41">Collar and Angioloni (2014)</xref>
</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>In addition to the high vitamin and mineral content, buckwheat differs from other foods by its high content of phenolic compounds, which were already found to have a positive effect on human health. The phenolics that can be found in buckwheat are well described in the section 1 of this review. The main responsible molecules of this biological activity in buckwheat are rutin and quercetin, both found in higher amounts in <italic>F. tataricum</italic> than in <italic>F. esculentum.</italic> These&#xa0;flavonols are correlated to antioxidant, anticancer, antihypertension, antibacterial and anti-inflammatory activities.</p>
<p>Rutin has demonstrated to be a potent antioxidant in several processes in human health. One of the most important functions of rutin is the ability to reduce oxidative stress and inflammatory responses by scavenging ROS or preventing their formation (<xref ref-type="bibr" rid="B148">Mehta et&#xa0;al., 2021</xref>), which is directly correlated with its antioxidant properties. Rutin, together with ascorbic acid, showed antioxidant, anti-inflammatory and antiapoptotic activities against UV-induced skin damage, suggesting that both compounds are potentially cytoprotective (<xref ref-type="bibr" rid="B74">G&#x119;gotek et&#xa0;al., 2019</xref>). This flavonoid is also a promising neuroprotective compound that may help to fight with neurodegenerative diseases by reducing proinflammatory cytokines and improving antioxidant enzyme activities, among other processes (<xref ref-type="bibr" rid="B62">Enogieru et&#xa0;al., 2018</xref>). This antioxidant activity is closely linked to their anti-cancer property. Rutin participates in the inhibition of proliferation, attenuation of superoxide production, and decrease adhesion and migration of human cancerous cells (<xref ref-type="bibr" rid="B194">Sghaier et&#xa0;al., 2016</xref>), showing ability to inhibit the proliferation of breast (<xref ref-type="bibr" rid="B241">Yang et&#xa0;al., 2017</xref>), colon (<xref ref-type="bibr" rid="B11">Alonso-Castro et&#xa0;al., 2013</xref>), renal (<xref ref-type="bibr" rid="B32">Caparica et&#xa0;al., 2020</xref>), lung (<xref ref-type="bibr" rid="B98">Imani et&#xa0;al., 2021</xref>), and prostate (<xref ref-type="bibr" rid="B54">Ding et&#xa0;al., 2017</xref>) cancer, as well as other tumors by the regulation of several signal pathways. Moreover, together with quercetin, rutin has the ability to prevent liver inflammatory injury (<xref ref-type="bibr" rid="B133">Lee et&#xa0;al., 2013</xref>), and antimicrobial synergistic effects with antibiotic activity against drug-resistant bacterial (<xref ref-type="bibr" rid="B10">Alnour et&#xa0;al., 2022</xref>).</p>
<p>Quercetin has also shown antidiabetic (<xref ref-type="bibr" rid="B179">Rezabakhsh et&#xa0;al., 2019</xref>), anti-inflammatory (<xref ref-type="bibr" rid="B174">Rafiq et&#xa0;al., 2015</xref>), antioxidant (<xref ref-type="bibr" rid="B134">Leyva-Soto et&#xa0;al., 2021</xref>), antimicrobial (<xref ref-type="bibr" rid="B160">Nguyen and Bhattacharya, 2022</xref>), anti-Alzheimer&#x2019;s (<xref ref-type="bibr" rid="B105">Joshi et&#xa0;al., 2022</xref>), and cardiovascular (<xref ref-type="bibr" rid="B44">Dabeek and Marra, 2019</xref>) activities in humans.</p>
<p>Phenolic acids, including hydroxybenzoic and hydroxycinnamic acids, are the specialized aromatic metabolites that confer typical organoleptic characteristics to foods. They are related to several human health benefits including antioxidant, anti-inflammatory (<xref ref-type="bibr" rid="B195">Shahidi and Zhong, 2015</xref>; <xref ref-type="bibr" rid="B220">Tresserra-Rimbau et&#xa0;al., 2018</xref>), immunoregulatory, anti-microbial, anti-thrombotic, cardioprotective, anti-cancer (<xref ref-type="bibr" rid="B14">Anantharaju et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B152">Mondal et&#xa0;al., 2021</xref>) and antidiabetic activities (<xref ref-type="bibr" rid="B124">Kumar and Goel, 2019</xref>). Phenolic acids can act delaying microbial growth and inhibiting lipid oxidation, prolonging food shelf-life (<xref ref-type="bibr" rid="B76">Galanakis, 2018</xref>). In deep, each phenolic acid can behave in a different way. For example, hydroxybenzoic acid could act as a mediator in colorectal cancer prevention (<xref ref-type="bibr" rid="B185">Sankaranarayanan et&#xa0;al., 2020</xref>) and as strong antioxidant potential (<xref ref-type="bibr" rid="B48">Das et&#xa0;al., 2022</xref>), while gallic acid shows hyperlipidemic, antihyperglycemic, cardioprotective and anti-cancer potential (<xref ref-type="bibr" rid="B244">Zanwar et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B205">Sourani et&#xa0;al., 2020</xref>), protocatechuic acid demonstrated antioxidant, antimicrobial, anti-inflammatory, antiapoptotic and antiproliferative activity (<xref ref-type="bibr" rid="B193">Semaming et&#xa0;al., 2014</xref>), vanillic acid has activity as hepatoprotective, cardioprotective, anti-apoptotic, anti-proliferative, and against DNA-induced damage (<xref ref-type="bibr" rid="B9">Almeida et&#xa0;al., 2016</xref>), caffeic acid has antioxidant and antibacterial properties (<xref ref-type="bibr" rid="B206">Spagnol et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B172">Possamai-Rossatto et&#xa0;al., 2021</xref>), and chlorogenic acid was reported to improve cognitive function in elders (<xref ref-type="bibr" rid="B111">Kato et&#xa0;al., 2018</xref>).</p>
<p>The high content of phenolic compounds in buckwheat makes it a highly nutritious and disease-protective crop.</p>
</sec>
<sec id="s7">
<label>7</label>
<title>Future areas of research</title>
<p>Future areas of research in buckwheat encompass various aspects of its cultivation, breeding, and utilization:</p>
<list list-type="simple">
<list-item>
<p>(i) <italic>Breeding for improved traits</italic>: research can focus on developing buckwheat varieties with enhanced traits such as disease resistance, tolerance to abiotic stress factors (e.g., drought, heat), improved nutritional content, and optimized agronomic characteristics, as increased pest control potential.</p>
</list-item>
<list-item>
<p>(ii) <italic>Molecular information</italic>: functional genomics could be applied to identify genes involved in biosynthesis of several identified phenolics, in order to achieve the isolation of these compounds.</p>
</list-item>
<list-item>
<p>(iii) <italic>Climate resilience</italic>: with changing climatic conditions, studying the adaptability of buckwheat to different environments and its response to drought, heat, and other climate-related stress factors can be valuable for future cultivation.</p>
</list-item>
<list-item>
<p>(iv) <italic>Utilization and value-added products</italic>: exploring the potential of buckwheat for applications such as functional foods, nutraceuticals, and industrial uses, can open up new avenues for its commercialization and utilization.</p>
</list-item>
<list-item>
<p>(v) <italic>Enhancing collaboration</italic> between farmers, weed scientists and plant breeders to advance in the breeding and exploitation of this crop, especially in agroecology systems for autonomous weed management.</p>
</list-item>
</list>
</sec>
<sec id="s8" sec-type="author-contributions">
<title>Author contributions</title>
<p>YV-&#xc1;: Conceptualization, Investigation, Writing &#x2013; original draft, Writing &#x2013; review and editing. MR: Funding acquisition, Resources, Supervision, Writing &#x2013; review and editing. AS-M: Project administration, Resources, Supervision, Writing &#x2013; review and editing.</p>
</sec>
</body>
<back>
<sec id="s9" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This work was supported by the European Union&#x2019;s Horizon 2020 research and innovation program under grant agreement no. 771367, ECOBREED, and under the Structural Grant of Xunta de Galicia with Grant number ED431C 2023/27.</p>
</sec>
<sec id="s10" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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